Document Type : Original Article

Authors

1 Ph.D Student, Department of Plant Production and Genetics, Faculty of Agriculture and Natural Resources, University of Mohaghegh Ardabili, Ardabil, Iran

2 Professor, Department of Plant Production and Genetics, Faculty of Agriculture and Natural Resources, University of Mohaghegh Ardabili, Ardabil, Iran

3 Associate Professor, Department of Plant Production and Genetics, Faculty of Agriculture and Natural Resources, University of Mohaghegh Ardabili, Ardabil, Iran

4 Associate Professor, Department of Water Engineering, Faculty of Agriculture and Natural Resources, University of Mohaghegh Ardabili, Ardabil, Iran

5 Dryland Agricultural Research Institute, Agricultural Research, Education and Extension Organization (AREEO), Gachsaran, Iran

Abstract

Introduction
Drought stress is one of the most important abiotic factors that can limit plant growth and yield. The response of plants to water limitation has been evaluated based on genetic, biochemical, and morpho-physiological traits. Plants are constantly affected by drought stress and re-irrigation. Therefore, rapid and efficient recovery from water deficit stress may be one of the key determinants of drought adaptation in plants. The aim of this research was the evaluation of drought stress tolerance and recovery in lentil cultivars after stress conditions.
Materials and methods
In order to evaluate the response of lentil cultivars to drought stress and re-irrigation, a factorial split-plot experiment based on a randomized complete block design with three replications was conducted in the greenhouse. Drought stress was applied at the flowering stage. The factors include 4 lentil cultivars (Namin landrace and Sepehr, Gachsaran, and Kimiya cultivars), drought stress (control (irrigation at 80% FC), medium stress (irrigation at 55% FC) and severe stress (irrigation at 30% FC)) and 3 sampling times (three and six days after drought and recovery (two days after re-irrigation)). All the plants were allowed to grow until the flowering stage (50 days after sowing) under well-watered conditions (80% FC (field capacity) of soil). Afterward, the plants were randomly assigned to three different groups and were exposed to different irrigation regimes including the control (well-watered and maintained at 80% FC), medium stress (watered and maintained at 55% FC), and severe drought stress (watered and maintained at 30% FC). The moisture content of the soil was controlled and maintained within a defined range using the weight method. Stress conditions were kept until the crop maturity and harvesting stage. The leaf samples from 5 seedlings of each pod were collected at 3 and 6 days after drought stress exposure, and two days after re-irrigation and used for physiological and biochemical analysis. The samples immediately were frozen in liquid nitrogen and stored at −80°C until analysis.
Results and discussion
The results showed that adaptation to drought stress was closely related to the recovery ability of plants. Drought stress caused a decrease in chlorophyll a, chlorophyll b, total chlorophyll, carotenoid, protein and proline, yield, and yield components. The reduction of these traits was more remarkable at six days after stress. However, during the recovery time remarkable increase was observed in these traits. The results showed that the correlation between H2O2 and MDA was significant and positive. Furthermore, drought stress increased the amount of H2O2 and MDA, which increased the activity of antioxidant enzymes (catalase, polyphenol oxidase, and peroxidase). An increase in the intensity and duration of the drought stress also caused an increase in proline (63%), H2O2, (19%), and MDA (110%) content, and the activity of CAT (33%), PP0 (56%), and POX (24%) compared to the control treatment. An increase in the intensity and duration of the drought stress also caused an increase in H2O2 and MDA content and the activity of antioxidant enzymes. In addition, in the recovery conditions, a significant reduction in the destructive effects of stress (H2O2, MDA content) and the activity of antioxidant enzymes was visible. The results of the present study indicated that the effects of drought stress on lentil cultivars' yield and yield components (seed numbers, number of pods, 100-seed weight, and seed yield) were varied. Drought stress at the flowering stage decreased the number of seeds (20%) and pods per plant (37%), and 100-seed weight (16%), which led to 29% yield losses. Although the Gachsaran cultivar had the highest yield under normal conditions. However, under drought stress conditions Gachsaran and Sepehr cultivars showed the highest plant yield. On the other hand, the Namin landrace exhibited the lowest yield (40%) under stress conditions.
Conclusion
The water stresses markedly increased the reactive oxygen species (ROS) level and impaired the biosynthesis of the photosynthetic pigment, resulting in the reduction of plant growth and yield with fewer seeds and pods number per plant. However, re-irrigation (recovery) remarkably improved plant growth and reduced the negative effects of drought stress, such as reducing the amount of MDA and H2O2 and improving the activity of antioxidant enzymes and proline content. In conclusion, based on physiological traits Gachsaran, and Sepehr cultivars seem to be suitable cultivars for culture in the regions challenged with water deficit stress.

Keywords

Main Subjects

 
Abrishamchi, P., Ganjeali, A., Sakeni, H., 2012. Evaluation of morphological traits,  proline content and antioxidant enzymes activity in chickpea genotypes (Cicer arietinum L.) under drought stress. Iranian Journal of Pulses Research. 3, 17-30. [In Persian with English summary]. https://doi.org/10.22067/IJPR.V1391I2.24695
Aebi, H., 1984. Catalase in vitro. Methods Enzymol. 105, 121–126. https://doi.org/10.1016/s0076-6879(84)05016-3
Ahmadi, A., Amini Dehaghi, M., Sedghi, M., Mansourifar, C., 2020. The effect of drought stress on antioxidant enzyme activity and chlorophyll content of some advanced genotypes of lentil (Lens culinaris Medik). Enviromental Stress in Crope Seinces. 12, 1105- 1116. https://doi.org/10.22077/escs.2019.1712.1390
Aranjuelo, I., Molero, G., Erice, G., Avice, J. C., Nogués, S., 2011. Plant physiology and proteomics reveal the leaf response to drought in alfalfa (Medicago sativa L.). Journal of Experimental Botany. 62, 111–123. https://doi.org/10.1093/jxb/erq249
Arnon, A., 1967. Method of extraction of chlorophyll in the plants. Agron. Journal. 23, 112–121.
Bahadoran, M., Abrishamchi, P., Ejtehadi, H., Ghassemzadeh, F.,2015. Study on some physiological characteristics of Salsola richteri in drought conditions in the two desert regions of the South Khorasan province. Plant Biology. 7, 1-14. [In Persian with English summary]. https://dorl.net/dor/20.1001.1.20088264.1394.7.24.2.0
Bates, L., Waldren, S., Teare, I., 1973. Rapid determination of free proline for water stress studies. Plant Soil. 39, 205–207. https://doi.org/10.1007/BF00018060
Blokhina, O., Virolainen E., Fagerstedt, K.V., 2003. Antioxidants, oxidative damage and oxygen deprivation stress: a review. Annals of Botany. 91, 179-194. https://doi.org/10.1093/aob/mcf118
Bradford, M.M., 1976. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Annals Biochem. 72, 248–254. https://doi.org/10.1016/0003-2697(76)90527-3
Chen, D., Wang, S., Cao, B., Cao, D., Leng, G., Li, H., Yin, L., Shan, L., Deng, X., 2016. Genotypic variation in growth and physiological response to drought stress and re-watering reveals the critical role of recovery in drought adaptation in maize seedlings. Frontiers in Plant Science. 6,1–15. https://doi.org/10.3389/fpls.2015.01241
Demiral, T., Türkan, I., 2004. Does exogenous glycine betaine affect antioxidative system of rice seedlings under NaCl treatment? Journal of Plant Physiology. 161, 1089-1100. https://doi.org/10.1016/j.jplph.2004.03.009
El Haddad, N., Rajendran, K., Smouni, A., Es-safi, N.E., Benbrahim, N., Mentag, R., Nayyar, H., 2020. Screening the FIGS set of lentil (Lens culinaris Medikus) germplasm for tolerance to terminal heat and combined drought-heat stress. Agronomy. 1–27. https://doi.org/10.3390/agronomy10071036
Ganjeali, A., Nezami, A., 2008. Ecophysiology and Limiting the Yield of Beans. Mashhad University Jihad Publishers, pp. 500. [In Persian].
Guo, Z., W. Ou, Lu S., Zhong. Q., 2006. Differential responses of antioxidative system to chilling and drought in four rice cultivars differing in sensitivity. Plant Physiology and Biochemistry. 44, 828-836. https://doi.org/10.1016/j.plaphy.2006.10.024
Hura, T., Grzesiak, S., Hura, K., Elisabeth Thiemt, E., Tokarz, K., Wedzony, M., 2007. Physiological and biochemical tools useful in drought-tolerance detection in genotypes of winter triticale: accumulation of ferulic acid correlates with drought tolerance. Annals of Botany. 100, 767–775. https://doi.org/10.1093/aob/mcm162
Kar, M., Mishra, D.,1976. Catalase, peroxidase and polyphenol oxidase activity during rice leaf senescence. Plant Physiology. 57, 315–319. https://doi.org/10.1104/pp.57.2.315
Kumar, S., Barpete, S., Kumar, J., Gupta, P., Sarker, A., 2013. Global Lentil Production: Constraints and Strategies. SATSA Mukhapatra–Annu Tech. 17, 1-13.
Ministry of Agricultural Jahad. 2021. Agricultural Products Statistics. p. 19. [In Persian].
Mollasadeghi, V., Dadbakhsh, A., 2011. Evaluation of some yield components in wheat genotypes under the influence of drought stress after flowering. Australian Journal of Basic and Applied Sciences. 5, 1137-1142.
Monakhova, O.F., Chemyadev II., 2002. Protective role of kartolin-4 in wheat plants exposed to soil drought. Applied Biochemistry and Microbiology. 38, 373–380. https://doi.org/10.1023/A:1016243424428
Muscolo, A., Sidari, M., Anastasi, U., Santonoceto, C., Maggio, A., 2013. Effect of PEG-induced drought stress on seed germination of four lentil genotypes. Journal of Plant Interactions. 9, 354–363. https://doi.org/10.1080/17429145.2013.835880
Pintó-Marijuan, M., Munné-Bosch, S., 2014. Photo-oxidative stress markers as a measure of abiotic stress-induced leaf senescence: advantages and limitations. Journal of Experimental Botany. 65, 3845–3857. https://doi.org/10.1093/jxb/eru086
Plaut, Z., Butow, B.J., Blumenthal, C.S., Wrigley, C.W., 2004. Transport of dry matter into meveloping wheat kernels and its contribution to grain yield under post-anthesis water deficit and elevated temperature. Field Crops Research. 86, 185–98. https://doi.org/10.1016/j.fcr.2003.08.005
Rahbarian, R., Khavari-nejad, R., Ganjeali, A., Bagheri A. R., Najafi, F., 2011. Drought stress effects on photosynthesis, chlorophyll fluorescence and water relations in tolerant and susceptible chickpea (Cicer arietinum L.) genotypes. Acta Biologica Cracoviensia. 53, 47-56. https://doi.org/10.2478/v10182-011-0007-2
Rasti Sani, M., Lahouti, M., Ganjeali, A., 2014. Effect of drought stress on some morphophysiological traits and chlorophyll fluorescence of red bean seedlings (Phaseolus vulgaris L.). Iranian Journal of Pulses Research. 5, 103-116. [In Persian with English summary]. https://doi.org/10.22067/IJPR.V1393I1.46210
Sabaghpour, S., Seyedi, F., Mahmoodi, A., Safikhani., M, Pezeshkpour, P., Rostemi, B., Kamel, M., Ferayedi, Y., Alahyar, N., Poursiabidi, M. 2013. Kimiya, a new high yielding lentil cultivar for moderate cold and semi warm climate of Iran. Seed and Plant Journal. 29, 397-399. [In Persian with English summary]. https://doi.org/10.22092/SPIJ.2017.111165
Sehgal, A., Sita, K, Bhandari, K., Kumar, S., Kumar, J., Vara Prasad, P.V., Siddique, K.H.M., Nayyar, H., 2019. Influence of drought and heat stress, applied independently or in combination during seed development, on qualitative and quantitative aspects of seeds of lentil (Lens culinaris Medikus) genotypes, differing in drought sensitivity. Plant Cell and Environment. 42, 198–211. https://doi.org/10.1111/pce.13328
Singh, D., Singh, CK., Taunk, J., Jadon, V., Pal, M., Gailkwad, K., 2019. Genome wide transcriptome analysis reveals vital role of heat responsive genes in regulatory mechanisms of lentil (Lens culinaris Medikus). Science Reporter. 9, 1–19. https://doi.org/10.1038/s41598-019-49496-0
Sinha, R., Kumar, A., Anil, P., Singh, K., 2018. Physiological, biochemical and molecular responses of lentil (Lens culinaris Medik) genotypes under drought stress. Indian Journal Plant Physiology. https://doi.org/10.1007/s40502-018-0411-7
Srivastava, R., Vasishtha, H., 2012. Saponins and lectins of Indian chickpeas (Cicer arietinum) and lentils (Lens culinaris). Indian Journal of Agricultural Biochemistry. 25, 44-47.
Talaat, NB., Shawky, BT., Ibrahim, AS., 2015. Alleviation of drought-induced oxidative stress in maize (Zea mays L.) plants by dual application of 24-epibrassinolide and spermine. Environmental and Experimental Botany. 113, 47–58. https://doi.org/10.1016/j.envexpbot.2015.01.006
Tariq, A., Pan, K., Olatunji, OA., Graciano, C., Li, Z., Wu, X., Chen, W., Song, D., Huang, D., Xue, T., Zhang, A., 2018. Phosphorous fertilization alleviates drought effects on Alnus cremastogyne by regulating its antioxidant and osmotic potential. Scientific Reports. 8, 1–11. https://doi.org/10.1038/s41598-018-24038-2
Valentovic, P., Luxova, M., Kolarovic, L., Gasparicova, O., 2006. Effect of osmotic stress on compatible solutes content, membrane stability and water relations in two maize cultivars. Plant, Soil and Environmen. 52, 186-191. https://doi.org/10.17221/3364-PSE
Ying, Y. Q., Song, L. L., Jacobs, D. F., Mei, L., Liu, P., Jin, S. H., et al., 2015. Physiological response to drought stress in Camptotheca acuminata seedlings from two provenances. Frontiers in Plant Science. 6, 361. https://doi.org/10.3389/fpls.2015.00361
Zaho, S., Xu, C., Zou, Q., 1994. Improvements of the method for measurement of malondialdehyde in plant tissue. Plant Physiology. Communications. 30, 207–210.
Zaragoza-Martínez, F., Lucho-Constantino, GG., Ponce-Noyola, T., Esparza-Garcıa, F., Poggi-Varaldo, H., Cerda-Garcı´a-Rojas, C.M., Trejo-Tapia, G., Ramous-Valdivia, C., 2016. Jasmonic acid stimulates the oxidative responses and triterpene production in Jatropha curcas cell suspension cultures through mevalonate as biosynthetic precursor. Plant Cell, Tissue and Organ Culture. 127, 47–56. https://doi.org/10.1007/s11240-016-1028-z